Recolonizing carnivores: Is cougar predation behaviorally mediated by bears?

Conservation and management efforts have resulted in population increases and range expansions for some apex predators, potentially changing trophic cascades and foraging behavior. Changes in sympatric carnivore and dominant scavenger populations provide opportunities to assess how carnivores affect one another. Cougars (Puma concolor) were the apex predator in the Great Basin of Nevada, USA, for over 80 years. Black bears (Ursus americanus) have recently recolonized the area and are known to heavily scavenge on cougar kills. To evaluate the impacts of sympatric, recolonizing bears on cougar foraging behavior in the Great Basin, we investigated kill sites of 31 cougars between 2009 and 2017 across a range of bear densities. We modeled the variation in feeding bout duration (number of nights spent feeding on a prey item) and the proportion of primary prey, mule deer (Odocoileus hemionus), in cougar diets using mixed‐effects models. We found that feeding bout duration was driven primarily by the size of the prey item being consumed, local bear density, and the presence of dependent kittens. The proportion of mule deer in cougar diet across all study areas declined over time, was lower for male cougars, increased with the presence of dependent kittens, and increased with higher bear densities. In sites with feral horses (Equus ferus), a novel large prey, cougar consumption of feral horses increased over time. Our results suggest that higher bear densities over time may reduce cougar feeding bout durations and influence the prey selection trade‐off for cougars when alternative, but more dangerous, large prey are available. Shifts in foraging behavior in multicarnivore systems can have cascading effects on prey selection. This study highlights the importance of measuring the impacts of sympatric apex predators and dominant scavengers on a shared resource base, providing a foundation for monitoring dynamic multipredator/scavenger systems.     

Anthropogenic Disturbance and Population Viability of Woodland Caribou in Ontario

One of the most challenging tasks in wildlife conservation and management is to clarify how spatial variation in land cover due to anthropogenic disturbance influences wildlife demography and long-term viability. To evaluate this, we compared rates of survival and population growth by woodland caribou (Rangifer tarandus caribou) from 2 study sites in northern Ontario, Canada that differed in the degree of anthropogenic disturbance because of commercial logging and road development, resulting in differences in predation risk due to gray wolves (Canis lupus). We used an individual-based model for population viability analysis (PVA) that incorporated adaptive patterns of caribou movement in relation to predation risk and food availability to predict stochastic variation in rates of caribou survival. Field estimates of annual survival rates for adult female caribou in the unlogged ( 0.90) and logged ( 0.76) study sites recorded during 2010–2014 did not differ significantly (P > 0.05) from values predicted by the individual-based PVA model (unlogged: = 0.87; logged: 0.79). Outcomes from the individual-based PVA model and a simpler stage-structured matrix model suggest that substantial differences in adult survival largely due to wolf predation are likely to lead to long-term decline of woodland caribou in the commercially logged landscape, whereas the unlogged landscape should be considerably more capable of sustaining caribou. Estimates of population growth rates (λ) for the 2010–2014 period differed little between the matrix model and the individual-based PVA model for the unlogged (matrix model = 1.01; individual-based model = 0.98) and logged landscape (matrix model = 0.88; individual-based model = 0.89). We applied the spatially explicit PVA model to assess the viability of woodland caribou across 14 woodland caribou ranges in Ontario. Outcomes of these simulations suggest that woodland caribou ranges that have experienced significant levels of commercial forestry activities in the past had annual growth rates <0.89, whereas caribou ranges that had not experienced commercial forestry operations had population growth rates >0.96. These differences were strongly related to regional variation in wolf densities. Our results suggest that increased wolf predation risk due to anthropogenic disturbance is of sufficient magnitude to cause appreciable risk of population decline in woodland caribou in Ontario. © 2020 The Authors. The Journal of Wildlife Management published by Wiley Periodicals, Inc. on behalf of The Wildlife Society.     

Cougar Prey Selection in a White-Tailed Deer and Mule Deer Community

Abstract Widespread mule deer (Odocoilus hemionous) declines coupled with white-tailed deer (O. virginianus) increases prompted us to investigate the role of cougar (Puma concolor) predation in a white-tailed deer, mule deer, and cougar community in northeast Washington, USA. We hypothesized that cougars select for and disproportionately prey on mule deer in such multiple-prey communities. We estimated relative annual and seasonal prey abundance (prey availability) and documented 60 cougar kills (prey usage) from 2002 to 2004. White-tailed deer and mule deer comprised 72% and 28% of the total large prey population and 60% and 40% of the total large prey killed, respectively. Cougars selected for mule deer on an annual basis (α_md = 0.63 vs. α_wt = 0.37; P = 0.066). We also detected strong seasonal selection for mule deer with cougars killing more mule deer in summer (α_md = 0.64) but not in winter (α_md = 0.53). Cougars showed no seasonal selection for white-tailed deer despite their higher relative abundance. The mean annual kill interval of 6.68 days between kills varied little by season (winter = 7.0 days/kill, summer = 6.6 days/kill; P = 0.78) or prey species (white-tailed deer = 7.0 days/kill, mule deer = 6.1 days/kill; P = 0.58). Kill locations for both prey species occurred at higher elevations during summer months (summer = 1,090 m, winter = 908 m; P = 0.066). We suspect that cougars are primarily subsisting on abundant white-tailed deer during winter but following these deer to higher elevations as they migrate to their summer ranges, resulting in a greater spatial overlap between cougars and mule deer and disproportionate predation on mule deer.     

Evidence for Depressed Reproduction of Golden Eagles in Washington

Beginning in 1977 the Washington Department of Fish and Wildlife conducted annual surveys to determine statewide golden eagle (Aquila chrysaetos) occupancy and productivity. Current interest in the regional and national status of the species prompted our investigation to determine utility of historical data in assessing trends in reproduction, and to test efficacy of a sampling protocol that surveyed randomly selected territories and also accounted for detection probability. We found evidence indicating poor reproduction from 38 annual surveys conducted at 301 known territories statewide between 1977 and 2014. At 256 territories in eastern Washington, USA, apparent occupancy was low ( = 50.9%) and nesting success declined by 22%. All reproductive parameters were higher than at 45 territories in western Washington. We tested efficacy of a sampling protocol in 2013 and 2014 by surveying 108 randomly selected eastern territories. Probability of detecting eagles for these years from ground (= 89%) was greater than from air (= 66%). Our estimate of territory occupancy, corrected by probability of detection, was lower in 2013 (= 56.7%, 95% CI = 46.3–66.7%) than in 2014 (= 73.7%, 95% CI = 64.8–81.7%), as was the estimated number of breeding pairs (2013: = 158, 95% CI = 151–164; 2014: = 187, 95% CI = 182–192). Higher productivity (young/occupied territory) in 2013 (= 0.59, 95% CI = 0.40–0.82) than in 2014 (= 0.41, 95% CI = 0.27–0.59) and lower proportions of ≥1 immature eagle among nesting pairs in 2013 (16%) than in 2014 (31%), suggested higher immature pairing among sampled pairs contributed to inter-year differences in these reproductive parameters. Current and historical evidence for depressed golden eagle nesting in Washington is consistent with documented effects from habitat conversion, prey declines, lead contamination, and wind power development. We recommend future surveys in eastern Washington adhere to the random sampling protocol and conduct surveys at regular intervals to allow for trend analysis of reproductive parameters to better monitor golden eagle status. Surveys in western Washington, conducted exclusively from ground at all nests, will improve detection and cost efficiency. © 2020 The Wildlife Society.     

Evaluating apparent competition in limiting the recovery of an endangered ungulate

Predation can disproportionately affect endangered prey populations when generalist predators are numerically linked to more abundant primary prey. Apparent competition, the term for this phenomenon, has been increasingly implicated in the declines of endangered prey populations. We examined the potential for apparent competition to limit the recovery of Sierra Nevada bighorn sheep (Ovis canadensis sierrae), an endangered subspecies under the US Endangered Species Act. Using a combination of location, demographic, and habitat data, we assessed whether cougar (Puma concolor) predation on endangered bighorn sheep was a consequence of their winter range overlap with abundant mule deer (Odocoileus hemionus). Consistent with the apparent competition hypothesis, bighorn sheep populations with higher spatial overlap with deer exhibited higher rates of cougar predation which had additive effects on adult survival. Bighorn sheep killed by cougars were primarily located within deer winter ranges, even though those areas constituted only a portion of the bighorn sheep winter ranges. We suspect that variation in sympatry between bighorn sheep and deer populations was largely driven by differences in habitat selection among bighorn sheep herds. Indeed, bighorn sheep herds that experienced the highest rates of predation and the greatest spatial overlap with deer also exhibited the strongest selection for low elevation habitat. Although predator-mediated apparent competition may limit some populations of bighorn sheep, it is not the primary factor limiting all populations, suggesting that the dynamics of different herds are highly idiosyncratic. Management plans for endangered species should consider the spatial distributions of key competitors and predators to reduce the potential for apparent competition to hijack conservation success.     

Cougar Kill Rate and Prey Composition in a Multiprey System

Abstract: Assessing the impact of large carnivores on ungulate prey has been challenging in part because even basic components of predation are difficult to measure. For cougars (Puma concolor), limited field data are available concerning fundamental aspects of predation, such as kill rate, or the influence of season, cougar demography, or prey vulnerability on predation, leading to uncertainty over how best to predict or interpret cougar-ungulate dynamics. Global Positioning System (GPS) telemetry used to locate predation events in the field is an efficient way to monitor large numbers of cougars over long periods in all seasons. We applied GPS telemetry techniques combined with occasional snow-tracking to locate 1,509 predation events for 53 marked and an unknown number of unmarked cougars and amassed 9,543 days of continuous predation monitoring for a subset of 42 GPS-collared cougars in west-central Alberta, Canada. Cougars killed ungulates at rates near the upper end of the previously recorded range, and demography substantially influenced annual kill rate in terms of both number of ungulates (subad F [SAF] = 24, subad M [SAM] = 31, ad M = 35, ad F = 42, ad F with kittens <6 months = 47, ad F with kittens <6 months = 67) and kg of prey (SAF = 1,441, SAM = 2,051, ad M = 4,708, ad F = 2,423, ad F with kittens <6 months = 2,794, ad F with kittens >6 months = 4,280). Demography also influenced prey composition; adult females subsisted primarily on deer (Odocoileus spp.), whereas adult males killed more large ungulates (e.g., moose [Alces alces]), and subadults incorporated the highest proportion of nonungulate prey. Predation patterns varied by season and cougars killed ungulates 1.5 times more frequently in summer when juveniles dominated the diet. Higher kill rate in summer appeared to be driven primarily by greater vulnerability of juvenile prey and secondarily by reduced handling time for smaller prey. Moreover, in accordance with predictions of the reproductive vulnerability hypothesis, female ungulates made up a higher proportion of cougar diet in spring just prior to and during the birthing period, whereas the proportion of males increased dramatically in autumn during the rut, supporting the notion that prey vulnerability influences cougar predation. Our results have implications for the impact cougars have on ungulate populations and have application for using cougar harvest to manage ungulates.     

Optimal Strategies for Managing Wildlife Harvest Under System Change

Wildlife populations are experiencing shifting dynamics due to climate and landscape change. Management policies that fail to account for non-stationary dynamics may fail to achieve management objectives. We establish a framework for understanding optimal strategies for managing a theoretical harvested population under non-stationarity. Building from harvest theory, we develop scenarios representing changes in population growth rate () or carrying capacity () and derive time-dependent optimal harvest policies using stochastic dynamic programming. We then evaluate the cost of falsely assuming stationarity by comparing the outcomes of forward projections in which either the optimal policy or a stationary policy is applied. When declines over time, the stationary policy leads to an underharvest of the population, resulting in less harvest over the short term but leaving the population in a higher-value state. When declines over time, the stationary policy leads to overharvest, resulting in greater harvest returns in the short term but leaving the population in a lower and potentially more vulnerable state. This work demonstrates the basic properties of time-dependent harvest management and provides a framework for evaluating the many outstanding questions about optimal management strategies under climate change. Published 2021. This article is a U.S. Government work and is in the public domain in the USA.     

Comparative Patterns of Predation by Cougars and Recolonizing Wolves in Montana's Madison Range

Abstract: Numerous studies have documented how prey may use antipredator strategies to reduce the risk of predation from a single predator. However, when a recolonizing predator enters an already complex predator—prey system, specific antipredator behaviors may conflict and avoidance of one predator may enhance vulnerability to another. We studied the patterns of prey selection by recolonizing wolves (Canis lupus) and cougars (Puma concolor) in response to prey resource selection in the northern Madison Range, Montana, USA. Elk (Cervus elaphus) were the primary prey for wolves, and mule deer (Odocoileus hemionus) were the primary prey for cougars, but elk made up an increasingly greater proportion of cougar kills annually. Although both predators preyed disproportionately on male elk, wolves were most likely to prey on males in poor physical condition. Although we found that the predators partitioned hunting habitats, structural complexity at wolf kill sites increased over time, whereas complexity of cougar kill sites decreased. We concluded that shifts by prey to structurally complex refugia were attempts by formerly naïve prey to lessen predation risk from wolves; nevertheless, shifting to more structurally complex refugia might have made prey more vulnerable to cougars. After a change in predator exposure, use of refugia may represent a compromise to minimize overall risk. As agencies formulate management strategies relative to wolf recolonization, the potential for interactive predation effects (i.e., facilitation or antagonism) should be considered.     

Disturbance-Mediated Apparent Competition Decouples in a Northern Boreal Caribou Range

The most widely reported threat to boreal and mountain populations of woodland caribou (Rangifer tarandus caribou; caribou) involves habitat- or disturbance-mediated apparent competition (DMAC). With DMAC, natural and anthropogenic disturbances that increase the abundance of deciduous-browsing cervids (e.g., moose [Alces alces], deer [Odocoileus spp.]) are thought to promote predator (especially wolf [Canis lupus]) numbers, which heightens predation risk to caribou. We know most about the effects of DMAC on caribou where the species is under threat by anthropogenic activities in relatively productive southern boreal and mountain systems. Yet, >60% of extant boreal caribou range in North America consists of northern shield and taiga ecoregions of low productivity where caribou may compete with only 1 ungulate species (moose) in the context of DMAC. In this environment, we know very little of how DMAC acts as a limiting factor to caribou. In Saskatchewan, Canada, from 2014–2018, using a combination of vegetation sampling, aerial surveys, and telemetry data (n = 38 wolves), we searched for evidence of DMAC (trends in data consistent with the hypothesis) in an 87,193-km² section of the Western Boreal Shield, a poorly productive but pristine region (0.18% of land cover classed as an anthropogenic feature) with a historically high fire-return interval (47% of stands aged <40 years). Despite the high levels of disturbance, moose density was relatively low (47 moose/1,000 km²), likely because of the scarcity of deciduous or mixed-wood stands and low abundance of deciduous browse in the young conifer stands that dominated the landscape. In contrast, boreal caribou density was relatively high for the species (37 caribou/1,000 km²). Wolf density (3.1 wolves/1,000 km²) and pack sizes ( = 4.0 wolves/pack) were low and resident (established) territories were large ( = 4,360 km²; 100% minimum convex polygon). The low density of wolves mirrored the low (standardized) ungulate biomass index (UBI; moose + boreal caribou) of the study area (0.36 UBI/km²). We conclude that wolf and hence caribou populations were not responding in accordance with the outcomes generally predicted by DMAC in our study area because the requisite strong, positive response to fire of deciduous-browse and alternate-prey abundance was lacking. As a limiting factor to caribou, DMAC is likely modulated at a macroecological scale by factors such as net primary productivity, a corollary to the general hypothesis that we advance here (i.e., primary productivity hypothesis of DMAC). We caution against managing for caribou based on the presumption of DMAC where the mechanism does not apply, which may include much of boreal caribou range in the north. © 2020 The Wildlife Society.     

Spatial Partitioning of Predation Risk in a Multiple Predator-Multiple Prey System

ABSTRACT Minimizing risk of predation from multiple predators can be difficult, particularly when the risk effects of one predator species may influence vulnerability to a second predator species. We decomposed spatial risk of predation in a 2-predator, 2-prey system into relative risk of encounter and, given an encounter, conditional relative risk of being killed. Then, we generated spatially explicit functions of total risk of predation for each prey species (elk [Cervus elaphus] and mule deer [Odocoileus hemionus]) by combining risks of encounter and kill. For both mule deer and elk, topographic and vegetation type effects, along with resource selection by their primary predator (cougars [Puma concolor] and wolves [Canis lupus], respectively), strongly influenced risk of encounter. Following an encounter, topographic and vegetation type effects altered the risk of predation for both ungulates. For mule deer, risk of direct predation was largely a function of cougar resource selection. However, for elk, risk of direct predation was not only a function of wolf occurrence, but also of habitat attributes that increased elk vulnerability to predation following an encounter. Our analysis of stage-based (i.e., encounter and kill) predation indicates that the risk effect of elk shifting to structurally complex habitat may ameliorate risk of direct predation by wolves but exacerbate risk of direct predation by cougars. Information on spatiotemporal patterns of predation will be become increasingly important as state agencies in the western United States face pressure to integrate predator and prey management.     

Eagle visitation rates to carrion in a winter scavenging guild

Understanding the behavioral ecology of species of conservation concern can help to inform better management. During winters 2011 through 2017, we placed camera traps at stations baited with carrion to investigate characteristics of winter scavenging by golden eagles (Aquila chrysaetos) and bald eagles (Haliaeetus leucocephalus) in eastern Washington and Oregon, USA. Our objectives were to better understand exposure risk of individual eagles to lead contaminants and evaluate factors that affect eagle visitation to carrion to inform measures that reduce lead exposure. We studied photo sequences from 108 traps ( = 2,725 ± 306 [SE] images/trap) and used plumage and physical characteristics to track visitation of 183 individual golden eagles and 90 bald eagles at deer (Odocoileus spp.) carrion until it was totally consumed. At least 1 eagle visited 76% of traps ( = 2.5 ± 0.3 unique eagles/trap). On average, an eagle visited a trap 3.4 ± 0.2 times (range = 1–19 visits) over 1.9 ± 0.1 days (range = 1–9 days). We used general linear mixed models to identify influences on number of eagle visits and pooled visit duration. Individual golden eagles visited carrion about 25% more often and 50% longer than bald eagles, and individual juvenile eagles visited carrion more often and longer than immature and adult eagles. On average, an eagle made an additional visit to carrion for every golden eagle that came to the same trap. Eagles spent less time at offal ( = 26.2 ± 6.4 min) than at a whole carcass ( = 92.9 ± 7.5 min), and understory vegetation immediately surrounding carrion was associated with a 30% reduction in visitation time. In the Pacific Northwest during winter, adult and juvenile golden eagles, by virtue of their abundance and visitation to carrion compared to the immature age class and bald eagles of all ages, have the highest potential for exposure to anthropogenic effects from carrion visitation. Concealment of offal piles in vegetation may reduce, but not eliminate, eagle use because of competing scavengers that expose carrion locations. We found no evidence that carrion proximity to nearest known nests, topography, or snow cover affect visitation by eagles. Thus, short of using alternative ammunition to lead, we recommend burial or removal of offal from hunter-killed ungulates. © 2019 The Wildlife Society.     

Genetic Characteristics of Restored Elk Populations in Kentucky

Translocations are a common management practice to restore or augment populations. Understanding the genetic consequences of translocation efforts is important for the long-term health of restored populations. The restoration of elk (Cervus canadensis) to Kentucky, USA, included source stocks from 6 western states, which were released at 8 sites in southeastern Kentucky during 1997–2002. We assessed genetic diversity in restored herds and compared genetic similarity to source stocks based on 15 microsatellite DNA loci. Genetic variation in the restored populations was comparable to source stocks ( allelic richness = 3.52 and 3.50; expected heterozygosity = 0.665 and 0.661 for restored and source, respectively). Genetic differentiation among all source and restored populations ranged from 0.000 to 0.065 for pairwise F_ST and 0.034 to 0.161 for pairwise Nei's D_A. Pairwise genetic differentiation and Bayesian clustering revealed that stocks from Utah and North Dakota, USA, contributed most to restored populations. Other western stocks appeared less successful and were not detected with our data, though our sampling was not exhaustive. We also inferred natural movements of elk among release sites by the presence of multiple genetic stocks. The success of the elk restoration effort in Kentucky may be due, in part, to the large number of elk (n = 1,548), repeated releases, and use of diverse source stocks. Future restoration efforts for elk in the eastern United States should consider the use of multiple stock sources and a large number of individuals. In addition, preservation of genetic samples of founder stock will enable detailed monitoring in the future. © 2020 The Authors. The Journal of Wildlife Management published by Wiley Periodicals, Inc. on behalf of The Wildlife Society.     

Bobcat Hair Cortisol Correlates with Land Use and Climate

Bobcats (Lynx rufus) have been increasing in abundance in the northeast United States despite a corresponding trend of increased anthropogenic land uses. Inhabiting areas of high human land use can affect stress levels, and hence cortisol titers, for wildlife species by increasing frequency of human interaction and altering habitats. In turn, increased cortisol levels can have negative effects at the individual and population level including decreased immune function, slowed growth and tissue repair, reduced reproductive capacity, and nutritional deficiencies. We quantified cortisol in bobcats across New Hampshire and Vermont, USA, using hair samples, then explored associations between hair cortisol and various organismal, land use, land cover, and climatic variables at 2 different spatial scales. Hair cortisol differed by season and bobcat mass. On average, cortisol levels were higher in fall than in spring, and larger bobcats had lower cortisol levels. Anthropogenic land uses—especially residential and agricultural uses—were the most important predictors of hair cortisol at the town scale ( area = 93 km²). At a larger scale (Wildlife Management Units; area = 1,256 km²), temperature and precipitation were better predictors of hair cortisol, suggesting that extreme weather may have significant effects on bobcat population dynamics. Our results highlight the importance of landscape composition and local conditions in the sustainable management of furbearer populations. © 2021 The Wildlife Society.     

Differential prey use by male and female cougars in Washington

Male and female predators are often assumed to have the same effects on prey. Because of differences in body size and behavior, however, male and female predators may use different species, sexes, and ages of prey, which could have important implications for wildlife conservation and management. We tested for differential prey use by male and female cougars (Puma concolor) from 2003 to 2008 in Washington State. We predicted that male cougars would kill a greater proportion of larger and older prey (i.e., adult elk [Cervus elaphus]), whereas females would kill smaller and younger prey (i.e., elk calves, mule deer [Odocoileus hemionus]). We marked cougars with Global Positioning System (GPS) radio collars and investigated 436 predation sites. We located prey remains at 345 sites from 9 male and 9 female cougars. We detected 184 mule deer, 142 elk, and 17 remains from 4 other species. We used log-linear modeling to detect differences in species and age of prey killed among cougar reproductive classes. Solitary females and females with dependent offspring killed more mule deer than elk (143 vs. 83, P < 0.01), whereas males killed more elk than mule deer (59 vs. 41, P < 0.01). Proportionately, males killed 4 times more adult elk than did females (24% vs. 6% of kills) and females killed 2 times more adult mule deer than did males (26% vs. 15% of kills). Managers should consider the effects of sex of predator in conservation and management of ungulates, particularly when managing for sensitive species. © 2011 The Wildlife Society.     

Influence of Landscape Characteristics on Hunter Space Use and Success

Sport hunting of ungulates is a predominant recreational pursuit and the primary tool for managing their populations in North America and beyond, given its influence on ungulate distributions, social organization, and population performance. Similarly, land management, such as motorized vehicle access, influences ungulate distributions during and outside hunting seasons. Although research on ungulate responses to hunting and land use is widespread, knowledge gaps persist about space use of hunters and what landscape features discriminate among hunt types and between successful and unsuccessful hunters. We used telemetry location data from hunters (n = 341) to estimate space use from 2008–2013 during 3 types of controlled, 5-day hunts for antlered mule deer (Odocoileus hemionus) and elk (Cervus canadensis) in northeastern Oregon, USA: archery elk, rifle deer, and rifle elk. To evaluate space use, we developed utilization distributions for each hunter, created core areas (50% contours) for groups of hunters, and derived several metrics of space-use overlap between successful and unsuccessful hunters. We also modeled predictors of space use using resource utilization functions with beta regression and stepwise model building. Hunter space use was compressed, with even the largest core area (unsuccessful rifle elk hunters) encompassing <16% (1,178 ha) of the area. We found strong similarities in space use of rifle hunters compared to archers, and core areas of successful hunters were markedly smaller than those of unsuccessful hunters (e.g., = 104 ha vs. 681 ha, respectively, for archers). Percentage cover and distance from open roads were the most consistent covariates in the 6 final models (successful vs. unsuccessful for each of 3 hunts) but with different signs. For example, predicted use of archery and rifle elk hunters increased with cover but decreased for rifle deer hunters. Although the same covariates were in the final models for unsuccessful and successful rifle elk hunters, their negligible spatial overlap suggested they sought those features in different locales, a pattern also documented for rifle deer hunters. Our models performed well (Spearman's rank correlation coefficients = 0.99 for 5 of 6 models), reflecting their utility for managing hunters and landscapes. Our results suggest that strategic management of open roads and forest cover can benefit managers seeking to balance hunter opportunity and satisfaction with harvest objectives, especially for species of special concern such as mule deer, and that differences in space use among hunter groups should be accounted for in hunting season designs. © 2021 The Wildlife Society. This article has been contributed to by US Government employees and their work is in the public domain in the USA.     

Seasonal Foraging Ecology of Non-Migratory Cougars in a System with Migrating Prey

We tested for seasonal differences in cougar (Puma concolor) foraging behaviors in the Southern Yellowstone Ecosystem, a multi-prey system in which ungulate prey migrate, and cougars do not. We recorded 411 winter prey and 239 summer prey killed by 28 female and 10 male cougars, and an additional 37 prey items by unmarked cougars. Deer composed 42.4% of summer cougar diets but only 7.2% of winter diets. Males and females, however, selected different proportions of different prey; male cougars selected more elk (Cervus elaphus) and moose (Alces alces) than females, while females killed greater proportions of bighorn sheep (Ovis canadensis), pronghorn (Antilocapra americana), mule deer (Odocoileus hemionus) and small prey than males. Kill rates did not vary by season or between males and females. In winter, cougars were more likely to kill prey on the landscape as: 1) elevation decreased, 2) distance to edge habitat decreased, 3) distance to large bodies of water decreased, and 4) steepness increased, whereas in summer, cougars were more likely to kill in areas as: 1) elevation decreased, 2) distance to edge habitat decreased, and 3) distance from large bodies of water increased. Our work highlighted that seasonal prey selection exhibited by stationary carnivores in systems with migratory prey is not only driven by changing prey vulnerability, but also by changing prey abundances. Elk and deer migrations may also be sustaining stationary cougar populations and creating apparent competition scenarios that result in higher predation rates on migratory bighorn sheep in winter and pronghorn in summer. Nevertheless, cougar predation on rare ungulates also appeared to be influenced by individual prey selection.     

Population genetics of Great Basin feral horses

The genetic make-up of Great Basin wild (feral) horses was investigated by blood typing studies. Blood samples of 975 feral horses from seven trap sites in Nevada and Oregon were tested by serological and electrophoretic techniques for genetic markers at 19 polymorphic loci. The average number of variants for the seven feral populations [72·1 ± 3·2 (SEM), range 62–85] was not significantly different from that of 16 domestic breeds (75·0 ± 11·5, range 58–105). The expected average frequency of heterozygotes per locus (average heterozygosity) for the feral populations (0·402 ± 0·009, range 0·368–0·442) was not significantly different from the domestic breeds (0·389 ± 0·045, range 0·295–0·443). Dendrograms constructed using pairwise comparisons of Nei's distance measurements substantiated anecdotal accounts of the origins of Great Basin horses from Iberian, American saddle horse and draft horse breeds.     

Long-Term Spatiotemporal Dynamics and Factors Associated with Trends in Bare-Nosed Wombats

Geographically widespread species present challenges for conservation assessment. We used long-term spotlight surveys to assess spatiotemporal dynamics of bare-nosed wombats (Vombatus ursinus), encompassing 34 years of surveys for the Tasmanian mainland sub-species (V. u. tasmaniensis, 1985–2018) and 25 years for the Flinders Island sub-species (V. u. ursinus, 1994–2018). Wombat populations increased on the Tasmanian mainland by 2.59 times and on Flinders Island by 3.51 times ( = 1.05 and 1.1 times increase/yr, respectively). At smaller spatial scales on mainland Tasmania, increases in wombat counts generally occurred within meteorological regions and regional zones, except for the Central North (West Tamar) region where a decrease in wombats is linked to a sarcoptic mange disease epizootic. We used generalized additive models to assess relationships between variables and wombat counts. The most supported variables at the mainland Tasmania scale were (in order of importance) year, positive associations with time-lagged minimum temperature, Tasmanian devil (Sarcophilus harrisii) counts, and moonlight, and a negative association with time-lagged rainfall. Among meteorological regions, variables associated with wombat counts exhibited some heterogeneity, with temperature and rainfall the most frequently associated variables. Our long-term, large-scale, and ecologically diverse analysis of bare-nosed wombats supports spotlight monitoring as a valuable, relatively simple, and affordable survey method in Tasmania and beyond. © 2021 The Wildlife Society.     

Ecological effects of fear: How spatiotemporal heterogeneity in predation risk influences mule deer access to forage in a sky‐island system

Forage availability and predation risk interact to affect habitat use of ungulates across many biomes. Within sky‐island habitats of the Mojave Desert, increased availability of diverse forage and cover may provide ungulates with unique opportunities to extend nutrient uptake and/or to mitigate predation risk. We addressed whether habitat use and foraging patterns of female mule deer (Odocoileus hemionus) responded to normalized difference vegetation index (NDVI), NDVI rate of change (green‐up), or the occurrence of cougars (Puma concolor). Female mule deer used available green‐up primarily in spring, although growing vegetation was available during other seasons. Mule deer and cougar shared similar habitat all year, and our models indicated cougars had a consistent, negative effect on mule deer access to growing vegetation, particularly in summer when cougar occurrence became concentrated at higher elevations. A seemingly late parturition date coincided with diminishing NDVI during the lactation period. Sky‐island populations, rarely studied, provide the opportunity to determine how mule deer respond to growing foliage along steep elevation and vegetation gradients when trapped with their predators and seasonally limited by aridity. Our findings indicate that fear of predation may restrict access to the forage resources found in sky islands.     

Distribution of Competition Potential Between Native Ungulates and Free-Roaming Equids on Western Rangelands

Free-roaming equids (i.e., feral horses [Equus caballus] and burros [Equus asinus]) are widely distributed and locally abundant across the rangelands of the western United States. The 1971 Wild Free Roaming Horse and Burro Act (WFRHBA) gave the Bureau of Land Management (BLM) and United States Forest Service (USFS) the legal authority to manage these animals on designated public lands. To fulfill this responsibility, federal agencies established an Appropriate Management Level (AML), defined as the number of horses or burros that can be sustained on a given management unit under prevailing environmental conditions and land uses. Although the WFRHBA specifies that feral equids must be managed in ecological balance with other land uses, including conservation of native wildlife, population control measures such as gathers, contraception, and adoptions have failed to keep pace with intrinsic growth rates. Over 80% of federally managed herds currently exceed prescribed population levels, making the potential for competition between native ungulates and feral equids a growing concern among state wildlife agencies. Mule deer (Odocoileus hemionus), pronghorn (Antilocapra americana), elk (Cervus canadensis), and bighorn sheep (Ovis canadensis) are of ecological and economic value to the states where they occur, and all exhibit some degree of distributional, habitat, or dietary overlap with horses or burros. Notwithstanding the scale of the problem, to date there have been no range-wide assessments of competition potential among native and feral ungulates for space, forage, or water. To address this need, we compiled demographic, jurisdictional, and species occurrence data collected from 2010–2019 by federal and state agencies. We used these data to map the distributions of 4 native ungulate species across federal equid management units (FEMUS) in 10 western states (n = 174). We then made within-state rankings of the 50 units that were ≥2 times over AML and encompassed ≥3 native ungulates. Collectively, FEMUs covered approximately 225,000 km², representing 18% of all BLM and USFS lands in affected states. Each FEMU supported ≥1 native ungulate and 14% contained all 4. The degree of overlap between native and feral species varied by state, ranging from <1% for mule deer in Montana, to 40% for bighorn sheep in Nevada. Oregon had the largest proportion of units that supported all 4 native ungulates (58%), whereas Montana and New Mexico had the fewest equids, but all populations were over target densities. Despite the perception that the problem of equid abundance is limited to the Great Basin states, high intrinsic growth rates and social constraints on management practices suggest all affected states should monitor range conditions and native ungulate demography in areas where forage and water resources are limited and expanding equid populations are a concern. © 2021 The Wildlife Society.